Spring-neap cycle as a major driver of temporal variations in feeding of intertidal fishes: Evidence from the sea catfish Sciades herzbergii (Ariidae) of equatorial west Atlantic mangrove creeks

On macrotidal coasts, short- to medium term variations in feeding of intertidal fishes are influenced by several interacting time scales. To identify the driver of major variations in the feeding habits of intertidal fish, we used the pemecou sea catfish Sciades herzbergii (Ariidae), an abundant intertidal benthic second order consumer, as a model species. We analyzed the influence of the spring-neap and the day-night cycle on intertidal abundance, stomach fullness, diet composition and food consumption of S. herzbergii using block nets set at slack high tides in two mangrove creeks in north Brazil. At spring tides, intertidal abundance, stomach fullness, and total daily consumption of S. herzbergii were on average 8.8, 1.9, and 3.8 times higher than at neap tides, respectively. At spring tides, Uca spp. and Grapsidae (mostly Pachygrapsus gracilis) dominated the diet, irrespective of the time of day. Other important food items were Insecta and the semiterrestrial crab Ucides cordatus. At neap tides, Capitellidae contributed to the diet of nightly inundations while no specimens were caught at daytime. Creek location had no effect on any variable. Results from our study area and evidence from other studies suggest that the spring-neap tide pulse is likely the major driver of short- to medium term variations in feeding of intertidal fishes. This has important implications for feeding-related issues on macro- and probably also on mesotidal coasts: (i) juvenile fishes may have fortnightly growth spurts, (ii) intertidal prey populations may suffer regular fluctuations in mortality, (iii) studies of the feeding ecology of intertidal fishes should cover the combined effects of the tidal, diel, lunar and seasonal cycles, and (iv) the modeling of food webs should consider the differences between highly dynamic spring tide and quieter neap tide conditions.     

Activity patterns, feeding and burrowing behaviour of the crab Ucides cordatus (Ucididae) in a high intertidal mangrove forest in North Brazil

Activity patterns, feeding and burrowing behaviour of the economically important semi-terrestrial mangrove crab Ucides cordatus (Ucididae, L. 1763) was studied in a high intertidal Rhizophora mangle forest stand in Bragança, North Brazil. Video observations in the rainy and dry season were conducted over 24 h cycles at different lunar phases to investigate the behaviour of these litter-feeding crabs outside their burrows. During the rainy season, crabs stayed inside their burrows for 79% and 92% of the time during day and night, respectively. Time spent for feeding, burrowing and other activities outside their burrows was significantly longer during the day with 9.9% (night: 1.7%) and at waning and waxing moon with 9% (full and new moon: 0.9%). At neap tides (no tidal inundation) foraging and feeding activities outside burrows were clearly light-dependent, increasing at dawn and decreasing at dusk. Highest activities during daytime relate to the visual localisation of food. During the dry season, crabs spent less time inside burrows at neap tides than during the rainy season (80% and 91%, respectively). However, time spent for feeding activities was similar during both seasons. During almost all observation periods crabs collected leaf litter, but rarely fed on it outside burrows. At neap tides nearly all available litter was collected, suggesting that the U. cordatus population is litter-limited during these times. At spring tides (regular tidal inundation) the surface activity of U. cordatus was tide-dependent. Crabs closed their burrow entrances 2-3 h before flooding and re-emerged as soon as the tide retreated. During the day, burrow maintenance was the second most frequent behaviour after feeding. Agonistic interactions were regularly observed and were mainly related to burrow defence. The mean foraging radius of the crabs was only 19 cm (max: 1 m) underneath high Rhizophora mangle trees where crab densities were high. The results point to a high competition for burrows and show that U. cordatus is territorial. It is concluded that several exogenous factors, in particular light, leaf litter availability, flooding of burrows and the presence of conspecifics are important in controlling the crabs' activity patterns.     

红树林生长带与潮汐水位关系的研究

根据海南东寨港林市村,廉江车扳镇那肋村,深圳锦锈中华等３个实验点１０条横断面红树林潮滩高程水准测量,林市村潮汐水位观测,潮汐特征水位及浸淹频率分布计算,参照有关港点潮汐实测或预测报资料,确定红树林外边界为平均海面或稍上,内边界（或林内最大高程）为回归潮平均高高潮位（或大潮平均高潮位）对国内外有关报道进行了评述,指出国内许多红树林文献中使用的“高潮位”,“中潮位”“低潮位”用潮汐学术语表述应为“回归     

Holocene reef evolution in a macrotidal setting: Buccaneer Archipelago,Kimberley Bioregion,Northwest Australia

This study uses information derived from cores to describe the Holocene accretion history of coral reefs in the macrotidal (up to 11 m tidal range) Buccaneer Archipelago of the southern Kimberley coast, Western Australia. The internal architecture of all cored reefs is broadly similar, constituting well-preserved detrital coral fragments, predominantly branching Acropora, in a poorly sorted sandy mud matrix. However, once the reefs reach sea level, they diverge into two types: low intertidal reefs that maintain their detrital character and develop relatively narrow, horizontal or gently sloping reef flats at approximately mean low water spring, and high intertidal reefs that develop broad coralline algal-dominated reef flats at elevations between mean low water neap and mean high water neap. The high intertidal reefs develop where strong, ebb-dominated, tidal asymmetry retains seawater over the low tide and allows continued accretion. Both reef types are ultimately constrained by sea level but differ in elevation by 3–4 m.     

Linkages between seagrass,mangrove and saltmarsh as fish habitat in the Botany Bay estuary,New South Wales

Shallow-water vegetated estuarine habitats, notably seagrass, mangrove and saltmarsh, are known to be important habitats for many species of small or juvenile fish in temperate Australia. However, the movement of fish between these habitats is poorly understood, and yet critical to the management of the estuarine fisheries resource. We installed a series of buoyant pop nets in adjacent stands of seagrass, mangrove and saltmarsh in order to determine how relative abundance of fishes varied through lunar cycles. Nets were released in all habitats at the peak of the monthly spring tide for 12 months, and in the seagrass habitat at the peak of the neap tide also. The assemblage of fish in each habitat differed during the spring tides. The seagrass assemblage differed between spring and neap tide, with the neap tide assemblage showing greater abundances of fish, particularly those species which visited the adjacent habitats when inundated during spring tides. The result supports the hypothesis that fish move from the seagrass to the adjacent mangrove and saltmarsh during spring tides, taking advantage of high abundances of zooplankton, and use seagrass as a refuge during lower tides. The restoration and preservation of mangrove and saltmarsh utility as fish habitat may in some situations be linked to the proximity of available seagrass.     

Mangrove inundation and nutrient dynamics from a GIS perspective

A digital elevation model describing topography, tide elevation and inundation degree and frequency of a mangrove forest in North Brazil is discussed in relation to existing phosphate and physicochemical data in waters of an adjacent tidal creek. Due to smooth topography, an increase of 20 cm in tidal height above average neap tides increases flooded area from about 50 to 80%. Analysis of the relationship between microtopography, tidal height and flooding rate showed that in the upper 60 cm of the mangrove forest, increases of 20 cm in topographical height resulted in a doubling of the inundation frequency. This can be particularly relevant for the analysis of nutrient mobilization and vegetation structure of infrequently inundated wetlands. Throughout the year, low-tide phosphate in creek water was inversely proportional to the maximum area flooded during high tide, this correlation being higher during the dry season. Similarly, the inverse relationship between flooded areas and low-tide/high-tide pH ratios was highly significant during the dry season and the beginning of the rainy season. Although the high correlations obtained are based on data pairs obtained at high and low tide, it has to clarified whether the association between inundation degree and creek water pH is relevant for the stability of P compounds in sediment on the short scale of a tidal cycle.     

The climbing behaviour of Cerithidea anticipata (Mollusca: Gastropoda): The roles of physical and biological factors

Abstract Many molluscs in tidal wetlands climb trees as the tide rises, a behaviour usually assumed to be a means of avoiding subtidal predators. Some species are more active during neap tides, when the access of subtidal predators to the forest is limited, but rest on trees during spring tides. Cerithidea anticipata, which inhabits the mangrove forests around Darwin Harbour (Northern Territory, Australia), displayed almost exactly the opposite pattern. This species climbed higher, and was less active, during neap tides that did not flood the forest than during spring tides. In experiments with tethered snails, individuals prevented from climbing died during neap tides, apparently from physiological stress. Further, individuals resting on trees around clearings, usually selected shaded sites. These results suggest that the major reason C. anticipata climbed was to avoid physiological stress during neap tides, not subtidal predators during spring tides. There was some evidence of predation under the canopy, but the rate was relatively low and the species responsible appeared to be resident in the forest.     

Feeding rhythms of the green-lipped mussel, Perna viridis (Linnaeus, 1758) (Bivalvia: Mytilidae) during spring and neap tidal cycles

Food availability and feeding responses of the green mussel Perna viridis were investigated for two complete tidal cycles during both spring and neap tides. Temporal changes in total particulate matter (TPM), particulate inorganic matter (PIM) and particulate organic matter (POM), were smaller during neap than spring tides. Seston characteristics at different times of a tidal cycle were compared for both spring and neap tides. Only during spring tides were TPM and PIM significantly higher at high tides while POM remained relatively constant (P>0.05). The clearance rate of the mussels underwent temporal variations with tides, and was a negative power function of TPM and a positive linear function of f (organic content), during both spring and neap tides. f was the key factor influencing filtration rate, organic ingestion rate, absorption rate and absorption efficiency. All feeding rates increased linearly with increases in organic content. Pseudofaeces were produced only during spring but not neap tides. Feeding rates and absorption efficiency were highest at low and lowest at high tides (P<0.01). There was no significant temporal change in the wet weight and protein content of the crystalline style with the tidal regime. For the digestive gland, alpha-amylase activity was higher at spring than at neap tides, and higher during high tides in a tidal cycle. The digestive gland cellulase activity did not change significantly with the tides. For the crystalline style, both the activity of cellulase and alpha-amylase were not significantly different (P>0.05) between spring and neap tides. Tidal rhythms in feeding and digestion in this species were likely controlled by temporal variations in food availability in the seawater. By adjusting feeding rates and enzymatic activities, absorption in Perna viridis remained constant, irrespective of the changes in food availability.     

Solute dynamics in a North Brazilian mangrove: the influence of sediment permeability and freshwater input

Although water in mangrove sediments influences nutrient cycling in both, mangrove forest and estuary, little information exists on seasonal and vertical distribution of dissolved organic and inorganic compounds in the sediment column. We studied the influence of sediment texture and chemistry, permeability (K), tides, and rainfall on dissolved organic carbon (DOC) and nitrogen (DON), dissolved inorganic phosphate (DIP) and salinity in creek and sediment waters of a mangrove in Pará, Brazil. Water samples were taken from boreholes and piezometers in the mangrove forest and from an adjacent tidal creek at neap and spring tides, during the dry and rainy season. Forest sediment was analysed for carbon (C), nitrogen (N), salinity and permeability. Clay, C and N decreased with depth. Sediment permeability (K) was lowest (<0.1 m day⁻¹) in the upper, clay-rich and crab-burrow-free mud layer. In the deeper, fine sand strata, K ranged from 0.7 to 1.8 m day⁻¹. Tidal range in the creek was 3.5 and 5.5 m for neap and spring tides, respectively. Salinity, DOC, DON and DIP in creek water were inversely related to tidal height. Piezometer data revealed significant water level changes in deeper, sandy sediment layer, which followed, time-lagged, the tidal fluctuations. In contrast, tide did not affect the water level in the upper sediment due to low permeability. Compared with creek water, sediment water was enriched in DOC, DON and DIP because of organic matter input and mineralization. In deeper layers, solute concentration was most likely affected by sorption processes (DOC and DIP) and reduction reactions (DIP). During the rainy season, DOC and DON in creek and sediment water were higher than in the dry season. DIP appeared invariant to seasonal changes. In the rainy season, salt flushing from surface sediments resulted in higher salinities at intermediate sediment depths, while in the deeper layers salinity was lower due to exchange with water from the tidal creek.     

Diel and tidal changes in the distribution and feeding habits of Japanese temperate bass Lateolabrax japonicus juveniles in the surf zone of Ariake Bay

Japanese temperate bass Lateolabrax japonicus juveniles recruit to the surf zone and grow by feeding on commonly occurring coastal copepods. However, little is known about diel and tidal patterns in their migration and feeding habits. We sampled wild juveniles during the neap and spring tides, over periods of 24 h, with small seine nets in the sand flat of the eastern part of Ariake Bay, Kyushu, western Japan. In both the neap and spring tides, abundance of juveniles significantly increased during the daytime, being highest around the time of high tide. The relative gut fullness indices of juveniles drastically increased in the morning (0700–0900) and during the flood tide in the daytime, while major prey copepod (Paracalanus spp.) density in the ambient water was relatively constant. We summarized that L. japonicus juveniles would migrate to the surf zone after sunrise to feed on copepods, and then emigrate from the surf zone after sunset. The migratory behavior of L. japonicus juveniles would be influenced by light (daytime) and feeding activity influenced by both light (morning) and tidal condition (flood tide). The intertidal region of the tidal flat was recognized to be one of the important habitats for L. japonicus during their early life history.     

Contribution to the feeding ecology of the banded puffer fish Colomesus psittacus (Tetraodontidae) in north Brazilian mangrove creeks.

Stomach contents were examined from 102 banded puffer, Colomesus psittacus (Tetraodontidae), caught from intertidal mangrove creeks at diurnal neap tides between June and September, 1997 (early dry season) near Bragan?a (north Brazil). The study found that C. psittacus were specialized predators of Cirripedia (Balanus spp.) and Brachyuran crabs (Uca spp., Pachygrapsus gracilis) (mean: 58 and 38% by dry weight, respectively), emphasizing a short food chain in the mangrove system. Cirripedia and Brachyura dominated the diet in all size classes, however, the prey spectrum narrowed with fish size. The mean daily consumption of Cirripedia and Brachyura was 6.2% body weight of C. psittacus. On average C. psittacus consumed 100.3 g x ha(-1) x d(-1) of Cirripedia and 178.7 g x ha(-1) x d(-1) of Brachyura (wet weight). The predation on Brachyuran crabs--a significant driver of fluxes of organic matter and energy in the system--provides C. psittacus with an important ecological function in the mangrove food web. A plant-animal interaction is proposed where C. psittacus exerts a mutually beneficial cleaning function on the Aufwuchs (Cirripedia and associated epibiota) of Rhizophora mangle stilt roots. Our results and those of other studies suggest that C. psittacus encounter optimum foraging conditions in the mangrove at high inundations at daylight (spring tide-day) whereas darkness and low inundations are linked to poor foraging conditions (neap tide-night). The C. psittacus resource could be used as an alternative income in the region in terms of i) sustainable catch and filet processing for exports to East Asia, ii) developing certified aquaculture methods for breeding puffers for the aquarium trade.     

Meiobenthos of hypersaline tropical mangrove sediment in relation to spring tide inundation

Tropical intertidal sediments often contain porewater of relatively high salinity, especially in areas exposed to longer periods without seawater inundation and high evaporation. Such an area exists on the west coast of Zanzibar: a high intertidal mangrove plateau, flooded only during spring high tides, with sediment porewater salinities commonly exceeding 100 ppt. A field survey was conducted in this area to examine variations in population density of major meiofaunal taxa and the assemblage structure of free-living marine nematodes during spring-neap tidal cycles. Samples were taken on seven occasions for two months, starting from the end of the rainy season. Porewater salinity remained high throughout the sampling period, ranging from 89 to 160 ppt. Neither spring tide inundation nor heavy rains lowered the salinity markedly. The meiofauna consisted only of four taxa, present on all sampling occasions: nematodes, harpacticoid copepods, plathyhelminthes and chironomids. Densities in surface sediments (0–5 cm) were low compared to other mangrove areas, ranging from 271 to 656 animals 10 cm^-2 with nematodes dominant on all sampling occasions (58–87%). Density fluctuations could not be explained by the effects of spring tide inundation, but the meiofauna showed significant correlations with grain size and organic material. Despite the wide range of salinity, only the numbers of chironomids were negatively correlated with increased salinity. Nematode species diversity was low in all samples, although altogether 28 species were recorded in the samples. Four species occurred in more than 50% of the samples (Microlaimus sp. (100%), Metalinhomoeus sp. (76%), Daptonema sp.l (56%), Chromadorina sp. (56%)) while 12 species were found only in one or two samples. Multidimensional scaling ordination (MDS) of the nematode species abundance data indicated little effects of spring tide inundation on the assemblage structure, but rather a successive change from wet to dry season with a reduction in species diversity and increased numbers of the dominant nematode species Microlaimus sp.     

High water roost selection by waders: maximizing feeding opportunities or avoiding predation?

During the highest spring tides the intertidal sediment flats of estuaries are fully inundated at high water, and waders have no choice but to move to supratidal roosts, e.g. on open farmland, saltpans or beaches. However, in many estuaries during the lowest neap or intermediate tides there are sectors of upper intertidal sediment flats that remain exposed even at the peak of high water, and so waders have the choice of roosting either there or in supratidal sites. In the Tagus Estuary, Portugal, as elsewhere, waders use both types of roosts during high water. Our main objective was to understand what makes waders opt for one of these two types of available roosts. We monitored wader use of saltpans and intertidal roosts from spring to neap tides, and measured foraging and alarm behaviour, prey availability and disturbance by predators. Most of the wader species studied chose intertidal (mudflat) roosts whenever these were available, and only roosted in saltpans during the peak of spring tides. We hypothesized that this preference was explained either by an attempt (i) to continue feeding into the high water period, or (ii) to minimize predation risk. Extending feeding time into the high water period did not seem to be very relevant for roost choice because both prey availability and foraging activity were low in both types of roosts. However, predator disturbance was several times higher in the saltpans than in the intertidal roosts, suggesting that this factor may be the determinant in the choice of roost type.     

Tidal movements of the macrofauna on an exposed sandy beach in South Africa

All epifaunal and shallow-burrowing, intertidal macrofauna on an exposed sandy beach appear to undergo tidal migrations which vary in extent but are all upshore on the incoming tide and downshore on the outgoing tide during day and night. In the crustaceans this is probably endogenous rhythmic behaviour and they show increased activity at night. In the molluscs this movement is not synchronised by an intrinsic mechanism but results from behavioural responses to changing physical conditions. The large wedge clam, Donax serra, shows no tidal migrations but does show a semilunar rhythm of movement from just above mean tide level (MTL) during springs to the low tide level during neap tides.     

Semilunar courtship rhythm of the fiddler crab<Emphasis Type="Italic"> Uca lactea</Emphasis> in a habitat with great tidal variation

Semilunar courtship rhythm is a widely distributed phenomenon among fiddler crabs in the genus Uca (Decapoda, Ocypodidae). Typically, synchronous courtship has been reported to peak near spring tides. To determine whether a region of large tidal variation shifts reproductive activity, we measured the frequency of specific courtship behaviors including claw-waving and semidome building for U. lactea males on Kanghwa Island, Korea. We found that synchronized courtship for U. lactea peaked near neap tides, whereas near the spring tides, seawater flooded the habitat and males predominantly fed on the mudflat. Although active females, which hold their burrows and usually feed on the mudflat, are abundant near to spring tides, males rarely claw-waved to attract females. This pattern is atypical for the species because other populations of U. lactea on Japan and Taiwan are synchronous around spring tides. We suggest that males invest most of their time in feeding during spring tides because foraging is limited during neap tides. During neap tides, males feed infrequently and thus expend stored energy on courtship signals. We conclude that patterns of reproductive synchrony may be dependent on food availability in periodically changing environments.     

Foraging strategy of the mudskipper Periophthalmus sobrinus Eggert in a Kenyan mangrove

The feeding behaviour of Periophthalmus sobrinus was studied in a population living along the channels of the Tana river delta. In November 1993 mudskippers were sampled every 2 tidal h for two consecutive spring and neap tides both during the day and night. The fishes were sexed and their stomach contents analysed. The results indicated that the mudskipper has a carnivorous diet and forages during both day and night although feeding greatly decreases after dark. Quantitative and qualitative differences in diet were found during the day and night of the two synodic and tidal phases. These differences were also demonstrable in the two sexes. The study showed that females of P. sobrinus are more selective than males and that differences between the two sexes may be related to reproductive condition. Furthermore, the mudskippers showed a tidal rhythm in feeding behaviour with peaks of activity around low tide (LT) on spring tides and around high tide (HT) on neap tides. Foraging activity seems to be influenced by the different spatial and temporal distributions of the mudskipper's prey caused by synodic, tidal and diel changes in the environment.     

Simulated sea level change alters anatomy, physiology, growth, and reproduction of red mangrove (Rhizophora mangle L.)

Tropical coastal forests – mangroves – will be one of the first ecosystems to be affected by altered sea levels accompanying global climate change. Responses of mangrove forests to changing sea levels depend on reactions of individual plants, yet such responses have not been addressed experimentally. We report data from a long-term greenhouse study that assessed physiological and individual growth responses of the dominant neotropical mangrove, Rhizophora mangle, to levels of inundation expected to occur in the Caribbean within 50–100 years. In this study, we grew potted plants in tanks with simulated semidiurnal (twice daily) high tides that approximated current conditions (MW plants), a 16-cm increase in sea level (LW plants), and a 16-cm decrease in sea level (HW plants). The experiment lasted 2½ years, beginning with mangrove seedlings and terminating after plants began to reproduce. Environmental (air temperature, relative humidity, photosynthetically active radiation) and edaphic conditions (pH, redox, soil sulfide) approximated field conditions in Belize, the source locale for the seedlings. HW plants were shorter and narrower, and produced fewer branches and leaves, responses correlated with the development of acid-sulfide soils in their pots. LW plants initially grew more rapidly than MW plants. However, the growth of LW plants slowed dramatically once they reached the sapling stage, and by the end of the experiment, MW plants were 10–20% larger in all measured growth parameters. Plants did not exhibit differences in allometric growth as a function of inundation. Anatomical characteristics of leaves did not differ among treatments. Both foliar C:N and root porosity decreased from LW through MW to HW. Relative to LW and HW plants, MW plants had 1–7% fewer stomata/mm², 6–21% greater maximum photosynthetic rates, 3–23% greater absolute relative growth rates (RGRs), and a 30% higher RGR for a given increase in net assimilation rate. Reduced growth of R. mangle under realistic conditions approximating future inundation depths likely will temper projected increased growth of this species under concomitant increases in the atmospheric concentration of CO₂.     

Field measurements and modeling of groundwater flow and biogeochemistry at Moses Hammock, a backbarrier island on the Georgia coast

A combination of field measurements, laboratory experiments and model simulations were used to characterize the groundwater biogeochemical dynamics along a shallow monitoring well transect on a coastal hammock. A switch in the redox status of the dissolved inorganic nitrogen (DIN) pool in the well at the upland/saltmarsh interface occurred over the spring-neap tidal transition: the DIN pool was dominated by nitrate during spring tide and by ammonium during neap tide. A density-dependent reaction-transport model was used to investigate the relative importance of individual processes to the observed N redox-switch. The observed N redox-switch was evaluated with regard to the roles of nitrification, denitrification, dissimilatory nitrate reduction to ammonium (DNRA), ammonium adsorption, and variations in inflowing water geochemistry between spring and neap tides. Transport was driven by measured pressure heads and process parameterizations were derived from field observations, targeted laboratory experiments, and the literature. Modeling results suggest that the variation in inflow water chemistry was the dominant driver of DIN dynamics and highlight the importance of spring-neap tide variations in the high marsh, which influences groundwater biogeochemistry at the marsh-upland transition.     

Temporal scales of variation in settlement and recruitment of the mussel Perna perna (Linnaeus, 1758)

Population dynamics of many intertidal organisms are strongly affected by the abundance and distribution of larvae arriving on the shore. In particular, not only absolute numbers of settlers but also the degree of synchronisation of settlement can have a strong influence on whether density-dependent or density-independent processes shape adult shape populations. Temporal variation in rates of settlement and recruitment of the mussel Perna perna on the south coast of South Africa was investigated using a nested spatial design at different temporal scales. Variability in settlement at spring tides was examined at two temporal scales: lunar (to investigate the effect of state of the moon on settlement) and tidal (to investigate the influence of state of the tide on mussel settlement). Recruitment over neap tides was examined at one temporal scale, fortnight (to investigate the effect of date on mussel recruitment).Strong temporal variation was evident for both settlement and recruitment, but not at all time scales. Distinct peaks of settler/recruit abundance were observed during the lunar and neap tide studies. Recruitment intensity differed over the course of the year, and pulsing of recruitment was generally synchronised among locations. However, the strength of pulsing differed dramatically among locations, giving a significant interaction between fortnight and location. The finest temporal scale, investigated in the tidal study, did not reveal a significant effect of the state of the tide on settlement. The state of the moon (new or full) was not significant as a main factor (p = 0.052), although generally more settlers arrived on the shore during new moon. Phase of the moon appeared to have an effect on settler abundances, but only when and where densities were high.     

Modulation of dissolved oxygen levels in a hypertidal estuary by sediment resuspension

Hyperconcentrated benthic layers, which form during neap tides, recruit much of the fine sediment population of the turbidity maximum of a hypertidal estuary. Measurements of tidal amplitude and suspended solids concentration reveal that resuspension of the hyperconcentrated layers occurs between three and eight tides after neap tides rather than during spring tides (12 to 15 tides after neaps). During these resuspension events, dissolved oxygen levels are reduced but recover by spring tides. Peak solids concentrations and critically depressed dissolved oxygen levels are out of phase with tidal current amplitude. Thus observations close to neap and spring tides do not necessaraly capture the extremes of the envelope of water quality conditions.